Trial Field Key to the Species of HYGROPHORUS in the Pacific Northwest

Notes on HEBELOMA in the Pacific Northwest Prepared for the Pacific Northwest Key Council By Ian Gibson, South Vancouver Island Mycological Society Copyright  2008, 2011 Pacific Northwest Key Council

INTRODUCTION

Considerably more field work must be done before a comprehensive key can be written for Hebeloma in the Pacific Northwest. It has been said in jest that there are only four species: veiled Hebeloma, poison pie (Hebeloma crustuliniforme), scaly-stalked Hebeloma (Hebeloma sinapizans group), and sweet-smelling Hebeloma (Hebeloma sacchariolens). That is just to illustrate the difficulty of distinguishing members of the genus which David Arora calls “another faceless and featureless collection of brownish mushrooms”.

A monograph for the veiled Hebelomas was written by A.H. Smith et al. in 1983, The veiled species of Hebeloma in the western United States. The monograph describes more than 90 species of which about a third had collections examined from the Pacific Northwest. Most of the rest were described from Colorado and could occur in the Pacific Northwest. The monograph has now been made available by the University of Michigan online at http://quod.lib.umich.edu/cgi/t/text/text-idx?c=fung1tc;cc=fung1tc;view=toc;idno=AAW6632.0001.001 The keys it contains should be consulted for detailed work. They have been abstracted below to contain only the species that Smith et al. document from the Pacific Northwest, in this case defined as British Columbia, Washington, Oregon, and Idaho. Identification is based on microscopic features.

Three other uncommon veil-less species are considered here under Subgenus Denudata. One of these, Hebeloma spoliatum, has been studied by Naohika Sagara and found to fruit only through the application of ammonia or ammonia-releasing nitrogenous material (such as rotting carcasses) to soil, thus it is among those Hebeloma species that could be a "corpse finder".

Descriptions of the veiled species are found in the monograph mentioned above. Hebeloma crustuliniforme, Hebeloma sacchariolens, and the Hebeloma sinapizans group are well described by Arora (1986). Microscopic details of Hebeloma crustuliniforme are in Smith (1949). Hebeloma incarnatulum is described in Smith (1984) and Trudell & Ammirati (2009). Hebeloma spoliatum is described in Smith(1938) and Smith (1984), Hebeloma arenosum by Burdsall et al.(1986), and Hebeloma avellaneum in Kauffman & Smith (1933) and Smith & Weber (1979). KEY TO SPECIES

1a Veil present, usually fibrillose to cortinate (Subgenus Hebeloma)...... 2 1b Veil absent (Subgenus Denudata)...... 501

2a Spores in profile view bean-shaped to elliptic or ovate, the apex +/- rounded (Section Mesophaea)...... 3 2b Spores +/- inequilateral in profile and narrowed to a blunt apex or apex +/- snoutlike (Section Hebeloma)...... 4

3a Spores 10-15 um long or longer (Subsection Subviscidae)...... 101 3b Spores 7-10 (11) um long (Subsection Mesophaeae)...... 201

4a (2b) Pileus white to pallid or cream color (Subsection Pallidae)...... Hebeloma salmonense 4b Pileus more highly colored when young than in above choice...... 5

5a When fresh the odor fragrant to pungent-aromatic (Subsection Praeolidae)...... Hebeloma praeolidum 5b Not as above (the odor if present +/- pungent to radishlike)...... 6

6a Spores 7-10(11) um long (Subsection Mesosporae)...... 301 6b Spores (9) 10-15 um or longer (Subsection Magnisporae)...... 401

101a (3a) Odor and taste of radish, lamellae pinkish gray, veil pallid...... Hebeloma vinaceogriseum 101b Odor and taste not distinctive...... 102

102a Odor and taste farinaceous, interior of stipe bay-red...... Hebeloma sterlingii 102b Odor and taste not distinctive, stipe not splitting lengthwise, gregarious not caespitose...... Hebeloma ollaliense

201a (3b) Cuticle of pileus an ixotrichodermium rarely collapsing to an ixolattice (study young pilei) (Stirps Pseudostrophosum)...... 202 201b Not as above (cuticle rarely an ixolattice in age)...... 203

202a Cheilocystidia 40-70 x 7-12 x 2.3-5 um...... Hebeloma pseudostrophosum 202b Cheilocystidia 28-43 x 5-7 x 7-9 um...... Hebeloma alpinicola

203a (201b) Pileus and/or gills staining dark brown to blackish on some basidiocarps in situ (Stirps Nigromaculatum)...... 204 203b Not staining as above...... 205

HEBELOMA - 2 - 204a Veil whitish (pallid)...... Hebeloma nigromaculatum 204b Veil (or at least outer layer) buff to pale tan...... Hebeloma angelesiense

205a (203b) Stipe not darkening at base in age (Stirps Pascuense)...... Hebeloma perigoense 205b Stipe soon darkening in basal area at least (Stirps Mesophaeum)...... 206

206a Odor and/or taste raphanoid...... 207 206b Odor and/or taste not as above...... Hebeloma mesophaeum

207a Spores 7-9 x 4.5-5.5 um, veil copious...... Hebeloma strophosum var. strophosum 207b Spores 8-11 x 5-6 um...... 208

208a Veil copious and remains long persistent on margin of pileus and/or stipe...... Hebeloma strophosum var. occidentale 208b Veil thin and scarcely leaving a zone of fibrils on the stipe...... Hebeloma mesophaeum

301a (6a) Stipe not darkening in lower part by maturity...... 302 301b Stipe darkening (often slowly) from the base upward...... 304

302a Taste bitter-farinaceous; pileus orange-brown when moist.Hebeloma aurantiellum 302b Not as above...... 303

303a Cheilocystidia 26-33 x 8-12 um...... Hebeloma immutabile 303b Cheilocystidia 40-67 x 4.5-7 um...... see Hebeloma fastibile

304a (301b) Spores 7-9 x 4-5 um...... Hebeloma olympianum 304b Spores larger...... 305

305a Odor fragrant; cheilocystidia 27-41 x 8-11 um...... Hebeloma pinetorum 305b Not as above...... 306

306a Cheilocystidia 18-26 x 3-4 um, many of them tibiiform...Hebeloma subhepaticum 306b Cheilocystidia not as above (at least wider than 3-4 um)...... Hebeloma parcivelum

401a (6b) Stipe not staining or discoloring in the lower portion by late maturity (Stirps Coniferarum)...... 402 401b Stipe soon darkening at base or lower portion, then upward...... 404

402a Cheilocystidia fusoid-ventricose, the apices subcapitate; odor pungent...... Hebeloma pungens 402b Not as above...... 403

403a Cheilocystidia (some of them) cylindric-subcapitate; pileus slimy viscid......

HEBELOMA - 3 - ...... Hebeloma kelloggense 403b Not as above...... Hebeloma pseudofastibile

404a (401b) Spores dextrinoid (medium to dark reddish brown in Melzer’s (Stirps Kuehneri)...... 405 404b Spores not dextrinoid, but in a mount a few may be found which become pale to +/- reddish brown in 30 minutes; (dried specimens are most reliable for this test) (Strips Oregonense)...... 408

405a Spores 9-12 um long...... 406 405b Spores 10-15 um or more long...... Hebeloma marginatulum

406a Odor and taste mild or odor weakly pungent...... 407 406b Odor and taste raphanoid...... Hebeloma pseudofastibile

407a Veil whitish...... Hebeloma occidentale 407b Veil grayish...... Hebeloma obscurum

408a (404b) Taste and usually the odor of the crushed context raphanoid...... 409 408b Not as above (odor +/- pungent in some and in others the taste farinaceous to bitter or at least not raphanoid)...... 413

409a Spores 9-12 um long...... 410 409b Spores (10) 12-15 um long...... 412

410a Spores 9-12 x 5-6.5 um...... Hebeloma fastibile 410b Spores 6.5-8.5 um wide...... 411

411a Veil remnants on stem white...... Hebeloma latisporum 411b Veil remnants pale ochraceous...... Hebeloma idahoense

412a (412b) Spores distinctly rough under a high-dry objective. .Hebeloma stanleyense 412b Spores appearing smooth under a high-dry objective or a 1.25 NA objective...... Hebeloma oregonense

413a (408b) Stipe staining yellow where injured...... Hebeloma lutescentipes 413b Stipe not staining yellow where handled...... Hebeloma oregonense

501a (1b) Strong odor sweet and aromatic, like burnt sugar, caramel, fruit candy, or orange blossom...... Hebeloma sacchariolens 501b Not as above, odor often radish-like...... 502

502a Pileus 4-13(20) cm across, typically brown to cinnamon, dark reddish brown, ocher-brown, or pinkish tan, often shaded with gray or overlaid with a pallid sheen toward margin, which is at first minutely cottony; stipe 1-3 cm thick, with

HEBELOMA - 4 - distinct pallid to brownish flakes or protruding scales (Arora) ...... Hebeloma sinapizans group 502b Not as above...... 503

503a Cap whitish to buff, pale tan, crust-colored, pinkish tan, or vinaceous brown, stipe 0.5- 1.5(2) cm thick, often with a dandruffy or scurfy apex but not normally with scales, radish-like odor...... Hebeloma crustuliniforme and Hebeloma incarnatulum

Hebeloma incarnatulum is apparently common in the Pacific Northwest and has been misidentified as H. crustuliniforme: H. incarnatulum is distinguished from H. crustuliniforme by lacking the conspicuous droplets on gill edges, having a basal bulb, having spores that turn dark reddish brown when mounted in Melzer's reagent, and having cheilocystidia that are narrow, not clavate, (Trudell & Ammirati 2009). Note however that descriptions of each species (Smith 1949, Smith 1984) include both cylindric and clavate cheilocystidia. Hebeloma incarnatulum is distinguished from other Hebelomas in Smith 1949 by vinaceous brown to pinkish tan cap, gills that do not become spotted and are seldom beaded with moisture, long white (non-discoloring) stem with bulbous base, absent veil, radish-like odor, bitterish to radish-like taste, and medium- sized, obscurely ornamented spores 9-12 x 5.5-7(7.5) microns [note also the preference for living Sphagnum moss].

503b Not as above...... 504

504a Pileus 3-10 cm across, ovate bell-shaped becoming expanded umbonate; pinkish gray to pecan brown; viscid, bald; gills crowded, narrow, +/- adnate; at first pinkish gray; edges white-floccose; stipe 5-10(12) cm x 0.6-1.5 cm, at times ovate-bulbous at base; +/- pruinose-mealy, becoming bald, basal mycelium white; cespitose to gregarious under conifers; spores 8-10 x 5-6 microns, pleurocystidia absent, cheilocystidia 30-80 x 5-8 microns, narrowly clavate to filamentose, agglutinated, clamp connections present, (Smith)...... Hebeloma avellaneum 504b Not as above...... 505

505a Restricted to conifer seedling nursing beds, stem narrowing gradually at base; spores (9.5)10-13(14) x 5-7 microns, ovoid, adaxially flattened, rugose [wrinkled] ...... Hebeloma arenosum 505b Not as above...... 506

506a Pileus bald, viscid, hygrophanous, vinaceous brown to tawny to buff; gills pallid becoming avellaneous or dull pinkish cinnamon; stipe appressed-silky, whitish to pallid in upper part and darker brown in lower part, often becoming twisted- striate; odor and taste mild...... Hebeloma spoliatum 506b Not as above...... many other undocumented or undescribed species

HEBELOMA - 5 - REFERENCES

Arora, David. 1986. Mushrooms Demystified. Ten Speed Press. Burdsall Jr., H.H., J.S. MacFall, M.A. Albers. 1986. “Hebeloma arenosa (Agaricales, Cortinariaceae), a new species from lake state nurseries” Mycologia 78(5): 861-865. Kauffman, C.H., A.H. Smith. 1933. “Agarics collected in the vicinity of Rock River, Michigan in 1929” Pap. Mich. Acad. Sci. Arts Lett. 17. 1933 Sagara, Naohiko. 1984. ‘On "Corpse Finder"’ McIlvainea 6:(2) 7-9. Smith, Alexander H. 1938. “New and Unusual Agarics from North America I.” Mycologia 30: 20-41. Smith, Alexander H. 1949 Mushrooms in Their Natural Habitats. University of Michigan Press, Ann Arbor. Smith, Alexander H., Smith Helen V., Weber, Nancy S. 1979. How to Know the Gilled Mushrooms. Wm. C. Brown Company, Dubuque, Iowa. Smith, Alexander H., V.S. Evenson, D.H. Mitchel. 1983. The Veiled Species of Hebeloma in the Western United States. University of Michigan Press, Ann Arbor. Smith, Alexander H. 1984. “Studies of Species of Hebeloma (Fr.) Kummer from the Great Lakes Region of North America I.” Sydowia 37:271-283. Trudell, Steve, Joe Ammirati. 2009. Mushrooms of the Pacific Northwest. Timber Press.

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